Person:

Smith, Tanya

The Aterian fossil hominins represent one of the most abundant series of human remains associated with Middle Stone Age/Middle Paleolithic assemblages in Africa. Their dates have been revised and they are now mostly assigned to a period between 90 and 35 ka. Although the Aterian human fossil record is exclusively Moroccan, Aterian assemblages are found throughout a vast geographical area extending to the Western Desert of Egypt. Their makers represent populations that were located close to the main gate to Eurasia and that immediately predated the last out-of-Africa exodus. In this chapter, we present an analysis of the Aterian dental remains. The sizes of the Aterian dentitions are particularly spectacular, especially for the post-canine dentition. This massiveness is reminiscent of the Middle Paleolithic modern humans from the Near East, but also of the early Homo sapiens in North and East Africa. Morphologically, this megadontia is expressed in the development of mass-additive traits. The Aterian dentition also displays relatively thick enamel. These features help to set some of the traits observed in Neandertals in perspective and highlight their primitive or derived nature. The Aterian morphological pattern is also important to consider when interpreting the dental morphology of the first modern humans in Eurasia.

Dental enamel thickness has received considerable attention in ecological models of the adaptive significance of primate morphology. Several authors have theorized that the degree of enamel thickness may reflect selective pressures related to the consumption of fallback foods (dietary items that may require complex processing and/or have low nutritional value) during times of preferred food scarcity. Others have speculated that enamel thickness reflects selection during mastication of foods with particular material properties (i.e., toughness and hardness). Orangutans prefer ripe fruit when available, but show interspecific and sex differences in the consumption of fallback foods (bark, leaves, and figs) and other preferred foods (certain seeds). Bornean orangutans (Pongo pygmaeus) have also been reported to masticate more mechanically demanding foods than Sumatran orangutans (Pongo abelii). To test these ecological models, we assessed two-dimensional enamel thickness in orangutan full dentitions using established histological and virtual quantification methods. No significant differences in average enamel thickness (AET) were found between species. We found significant differences in the components of enamel thickness indices between sexes, with males showing greater enamel-dentine junction lengths and dentine core areas, and thus relatively thinner enamel than females. Comparisons of individuals of known sex and species revealed a dentition-wide trend for Bornean females to show greater AET than Sumatran females. Differences between small samples of males were less evident. These data provide only limited support for ecological explanations of enamel thickness patterns within great ape genera. Future studies of dietary ecology and enamel thickness should consider sex differences more systematically.

Humans have an unusual life history, with an early weaning age, long childhood, late first reproduction, short interbirth intervals, and long lifespan. In contrast, great apes wean later, reproduce earlier, and have longer intervals between births. Despite 80 y of speculation, the origins of these developmental patterns in Homo sapiens remain unknown. Because they record daily growth during formation, teeth provide important insights, revealing that australopithecines and early Homo had more rapid ontogenies than recent humans. Dental development in later Homo species has been intensely debated, most notably the issue of whether Neanderthals and H. sapiens differ. Here we apply synchrotron virtual histology to a geographically and temporally diverse sample of Middle Paleolithic juveniles, including Neanderthals, to assess tooth formation and calculate age at death from dental microstructure. We find that most Neanderthal tooth crowns grew more rapidly than modern human teeth, resulting in significantly faster dental maturation. In contrast, Middle Paleolithic H. sapiens juveniles show greater similarity to recent humans. These findings are consistent with recent cranial and molecular evidence for subtle developmental differences between Neanderthals and H. sapiens. When compared with earlier hominin taxa, both Neanderthals and H. sapiens have extended the duration of dental development. This period of dental immaturity is particularly prolonged in modern humans.

Dental enamel thickness continues to feature prominently in anthropological studies of ape and human evolution, as well as studies of preventative oral care and treatment. Traditional studies of enamel thickness require physical sectioning of teeth for linear and scaled measurements. Recent applications of microtomographic imaging allow scientists to employ larger and more diverse samples, including global samples of recent humans as well as fossil hominin teeth. Unfortunately, little is known about the degree of enamel thickness variation among human populations, particularly across the dentition. This study employed microtomography to virtually image, section, and quantify the average enamel thickness of a sample of clinically extracted Indonesian canine and premolar teeth. This virtual sample was compared to physically sectioned African and European teeth. The results demonstrate that average enamel thickness is similar among human dentitions; no significant differ- ences were detected within tooth positions, which is surprising given developmental differences be- tween European and African canines and premolars. When populations were combined, differences were found in average enamel thickness between maxillary and mandibular premolars, and between canines and premolars within both dental arcades. This finding is potentially due to differences in pre- molar morphology and a trend of increasing enamel thickness distally throughout the dentition. The finding of limited population variation within tooth positions and significant variation between tooth positions is consistent with previous two-dimensional and three-dimensional studies of human molar enamel thickness. Average enamel thickness in canines and premolars does not differ between the sexes in our sample, although male teeth tend to have larger enamel and dentine cross-sectional areas, enamel–dentine junction lengths, and bi-cervical diameters. Males have significantly greater dentine area and enamel–dentine junction length than females for maxillary canines and premolars. The results of this study suggest that enamel thickness values in mixed-populations of humans are appropriate for comparisons with fossil hominins.

The Thomas Quarry I locality was made famous in 1969 with the discovery of a human half-mandible in a cave. In 1985, further investigations revealed the presence of a Lower Acheulean assemblage in lower units of the section. From 1988 onwards, modern controlled excavations took place within the framework of the Franco-Moroccan co-operative project “Casablanca”. Acheulean artefacts, a rich mammalian fauna and four hominid teeth have been excavated from the cave. The faunal set indicates an open woodland environment. Carcasses were processed by carnivores, but cut-marks are absent, which raises the question of any human role in the bone accumulations. Stone knapping was mainly oriented towards flake production and a few bifaces have been imported into the site. Laser ablation ICP-MS dating combining the ESR and U-series data for the modelling of the U-uptake has given an US/ESR age of 501+94−76 ka for a human premolar while new OSL measurements yielded an age of 420 ± 34 ka for sediments immediately above the dated tooth and 391 ± 32 ka below. Nevertheless, biostratigraphy and lithostratigraphy point towards a greater antiquity.

Developmental studies consistently suggest that teeth are more buffered from the environment than other skeletal elements. The surprising finding of late tooth eruption in wild chimpanzees (Zihlman et al., 2004) warrants reassessment in a broader study of crown and root formation. Here we re-examine the skeletal collection of Taï Forest juvenile chimpanzees using radiography and physical examination. Several new individuals are included, along with genetic and histological assessments of questionable identities. Only half of the Taï juveniles employed by Zihlman et al. (2004) have age of death known with accuracy sufficient for precise comparisons with captive chimpanzees. One key individual in the former study, misidentified during field recovery as Xindra (age 8.3), is re-identified as Goshu (age 6.4). For crown formation we find that onset and duration greatly overlap captive chimpanzees, whereas root development may be more susceptible to acceleration in captive individuals. Kuykendall's (1996) equation relating captive tooth formation stage to age gives reasonable estimates of young wild subjects' true ages. Direct comparisons of tooth eruption ages are limited. A key 3.76 year-old individual likely possessed an emerging mandibular M1 at death (previously estimated from the maxillary molar as occurring at 4.1 years). Wild individuals appear to fall near the middle or latter half of captive eruption ranges. While minor developmental differences are apparent in some comparisons, our reanalysis does not show an “unambiguous pattern” of slower tooth formation in this wild environment. These data do not undermine recent developmental studies of the comparative life histories of fossil hominins.

Analyses of hominin dental remains conventionally include measurements of tooth crown sizes and descriptions of occlusal morphology such as minor accessory cusps, fissure patterns, and ridges (e.g., Wood, 1981, Aiello and Dean, 1990 and Bailey, 2006). Following Dahlberg, 1951 and Turner et al., 1991 developed a formal system for dividing these ‘non-metric’ aspects of dental morphology into discrete categories. This system, termed the Arizona State University Dental Anthropology System (ASUDAS), is an effective tool for discriminating among modern human populations and for assessing inter-population relationships (Scott and Turner, 1997). Other researchers later used ASUDAS to examine the evolutionary relationships of various Pleistocene hominins (Irish and Guatelli-Steinberg, 2003 and Martinón-Torres et al., 2007; but see Bailey et al., 2009). Most recently, Irish et al. (2013) used ASUDAS to assess the phylogenetic position of Australopithecus sediba, finding support for both an Au. sediba + Au. africanus clade and a clade uniting South African australopiths with Homo. However, as Kimbel (2013) has argued, there are theoretical issues with applying ASUDAS to assess phylogenetic relationships from small samples of fossil hominin dental remains. Here we explore the suitability of applying a method developed for partitioning among modern human populations to assess interspecies relationships among fossil hominins. We then discuss the ramifications of different choices made during phylogenetic estimation, including those pertaining to character weighting, clade support, and outgroup composition. We find that slight alteration of phylogenetic assumptions leads to numerous equally possible evolutionary reconstructions for Au. sediba.

Orangutans (Pongo) are the only great ape genus with a substantial Pleistocene and Holocene fossil record, demonstrating a much larger geographic range than extant populations. In addition to having an extensive fossil record, Pongo shows several convergent morphological similarities with Homo, including a trend of dental reduction during the past million years. While studies have documented variation in dental tissue proportions among species of Homo, little is known about variation in enamel thickness within fossil orangutans. Here we assess dental tissue proportions, including conventional enamel thickness indices, in a large sample of fossil orangutan postcanine teeth from mainland Asia and Indonesia. We find few differences between regions, except for significantly lower average enamel thickness (AET) values in Indonesian mandibular first molars. Differences between fossil and extant orangutans are more marked, with fossil Pongo showing higher AET in most postcanine teeth. These differences are significant for maxillary and mandibular first molars. Fossil orangutans show higher AET than extant Pongo due to greater enamel cap areas, which exceed increases in enamel-dentine junction length (due to geometric scaling of areas and lengths for the AET index calculation). We also find greater dentine areas in fossil orangutans, but relative enamel thickness indices do not differ between fossil and extant taxa. When changes in dental tissue proportions between fossil and extant orangutans are compared with fossil and recent Homo sapiens, Pongo appears to show isometric reduction in enamel and dentine, while crown reduction in H. sapiens appears to be due to preferential loss of dentine. Disparate selective pressures or developmental constraints may underlie these patterns. Finally, the finding of moderately thick molar enamel in fossil orangutans may represent an additional convergent dental similarity with Homo erectus, complicating attempts to distinguish these taxa in mixed Asian faunas.

A comparative study has been made of human and great ape molar tooth enamel. Nanoindentation techniques are used to map profiles of elastic modulus and hardness across sections from the enamel–dentin junction to the outer tooth surface. The measured data profiles overlap between species, suggesting a degree of commonality in material properties. Using established deformation and fracture relations, critical loads to produce function-threatening damage in the enamel of each species are calculated for characteristic tooth sizes and enamel thicknesses. The results suggest that differences in load-bearing capacity of molar teeth in primates are less a function of underlying material properties than of morphology.